charles wright and cuban palms. 1. resurrection

PALMS Moya López & Méndez Santos: Coccothrinax Vol. 62(1) 2018

42

The palm family has undergone an importantprocess of diversification in Cuba, resulting in15 genera with 79 species, 8 hybrids and 11infraspecific taxa (Moya & Leyva 2000,updated by the first author). However, recentpublications (Suárez 2015, Verdecia 2016,Moya et al. 2017), suggest that the taxonomicrichness of the group in Cuba is not yet fullyknown.

This abundance of palms in Cuba has attracteda great deal of attention. Numerous researchersfrom various countries have made contri-

butions, but they have not always taken intoaccount the historical record. Therefore, forthe development of a robust taxonomy ofCuban palms today, the task of reanalyzinghistorical information becomes a priority, inorder to correct errors and eliminate possiblecauses of ambiguity and confusion.

In the genus Coccothrinax, restricted withinthe Caribbean Basin, 55 species have beendescribed, seven infraspecific taxa and onehybrid. Cuba is its center of diversity where 39species are found (38 of them endemic), seven

Charles Wrightand CubanPalms. 1.Resurrectionof Coccothrinaxacuminata

CELIO E. MOYA LÓPEZSección de Palmas,Sociedad Cubana deBotánica 90 South Blvd. Apt. 2C, Boynton Beach, Florida33435 [email protected]

AND

ISIDRO E. MÉNDEZ SANTOSUniversidad de Camagüey“Ignacio AgramonteLoynaz”Carretera deCircunvalación Norte, Km5 ½, Camagüey, [email protected]

A new taxonomic treatment to demonstrate the legitimacy of the name

Coccothrinax acuminata Becc. (Arecaceae) is presented. Two taxa, Coccothrinax

miraguama var. novo-geronensis and C. miraguama subsp. arenicola, are treated

as new synonyms. Three lectotypes are designated, and 38 isolectotypes not

mentioned in previous publications are indicated.

PALMS 62(1): 42–50


43

1. Stem with leaf sheaths and older infrutescence of Coccothrinax acuminata in Cortés, Pinar del Río. Photo byC.E. Moya.

subspecies and one hybrid. Nine species havebeen reported in Hispaniola (seven of themendemic). Of the remaining, four are limitedto the islands, two share their distributionbetween the islands and continental territoryand only one is confined to the mainland(Jestrow et al. 2017).

The taxonomic treatment of this genus cannotbe considered complete. On the one hand,new descriptions continue to be published;three occurred in the last four years (Mejía &García 2013, Suárez 2015 and Moya et al 2017).On the other hand, it is still necessary to refinepart of the nomenclature used in previousaccounts of currently recognized taxa.

A particularly complex taxonomic andnomenclatural situation occurs in westernCuba, around what could preliminarily becalled the “Coccothrinax acuminata complex,”a name used by different authors, althoughnot always legitimately. It is a group that growsin the southern parts of the provinces of Pinardel Río and Artemisa, as well as in central andnorthern Isla de la Juventud. Taxa in thiscomplex are characterized by the presence ofshort, pendulous mature infructescences, witha rachis partially down-curved, leaves that are5/4 orbicular, with segments 12–15 cm longmeasured from the “shoulder” to the apex,thin leaf sheath strands 0.5–1 mm wide anddensely woven in three layers (Fig. 1). In

addition to the main species, two otherinfraspecific taxa, Coccothrinax miraguama(novo-geronensis) Becc. and C. miraguama subsp.arenicola (León) Borhidi & O.Muñiz, areinvolved.

The objectives of this paper are to offer a newtaxonomic system that better reflects thenatural variability in this complex, to evaluatethe nomenclature used historically for it, todetermine the accepted name, to reveal thelocation of the type material in differentherbaria and to order the synonymy.

Materials and Methods

The nomenclature and taxonomy of what werefer to in this paper as “Coccothrinax acuminatacomplex” was investigated. A review was doneof the protologues of the names used bydifferent authors and of the differentdescriptions available in the main treatments.Expedition notes and species catalogs were alsostudied. A taxonomic study was made from areview of herbarium specimens and fieldstudies conducted by the first author for over30 years.

The following works were consulted: Sauvalle(1871, 1873), Gómez de la Maza (1893),Sargent (1899), Schumann (1901), Beccari(1907, 1913), Britton (1910, 1916), Shafer(1913), Jennings (1917), León (1918), Dahlgren(1936), León (1939, 1946), Borhidi and Muñiz


44

2. Coccothrinax acuminata growing together with Acoelorrhaphe wrightii (left and background) in Cortés.Photo by C.E. Moya.

(1971), Glassman (1972), Muñiz and Borhidi(1982), Chiappy et al. (1986), Cejas andHerrera (1995), Hernández et al. (1995), Moyaand Leiva (2000), Govaerts and Dransfield(2005), Govaerts et al. (2011), AcevedoRodríguez and Strong (2012), Novo et al.(2015) and Greuter and Rankin (2016). Thematerials used by Beccari were reviewed inCuccuini and Nepi (2006). The effectiveness,validity and legitimacy of each name used waschecked using the International Code ofNomenclature for algae, fungi, and plants(ICN) (McNeill et al. 2012).

128 specimens of 38 collections (including 45types) were reviewed from the followingherbaria: BRU, CM, F, FI, GH, HAC, HCM, K,LE, M, MO, NY, P, US (acronyms sensu Thiers,2016). Special attention was paid to the studyof specimen duplicates (Wright 3966) in sevenof these herbaria, as well as clarification of thelocation of the lectotypes and isolectotypes. Astudy was made of the exact locations of thecollections made by Charles Wright that arebasic to an understanding of the taxonomyand nomenclature of the taxon, as was a studydone of the contrasting data annotated by himon herbarium labels. We also reviewedinformation available in the literature(Underwood 1905, Howard 1988) and thecurrent cartography and phytogeography ofthe region (Borhidi 1996).

Field expeditions were carried out in localitieswhere the presence of the taxon was knownin the province of Pinar del Río: north ofCortés (Fig. 2), municipality Sandino; nearHerradura, municipality Consolación del Sur;savannahs of El Sábalo, municipality Guanes,as well as the area surrounding La Cañada, LaFe, Siguanea, Las Nuevas, Hotel Colony andNueva Gerona of Isla de la Juventud. Alsovisited were the localities of Las Pozas,Cajalbana and Viñales in western Cuba, whereother species of the genus are reported.Diagnostic characters were studied in situ, anda preliminary key was developed todifferentiate it from the other Coccothrinaxspecies in that part of the country.

Results and Discussion

The evaluation and comparison of morpho-logical, phenological and phytogeographicevidence, from plants in situ (during field trips)and from 126 herbarium specimens ofCoccothrinax collected in western Cuba led themain author to the preliminary conclusionthat all specimens previously designated as

C. acuminata, C. miraguama (novo-geronensis)and C. miraguama subsp. arenicola constitutea single taxon.

Taxonomic and nomenclatural history

The first event relevant to the group understudy was the collection of Coccothrinaxacuminata made by Charles Wright and giventhe number 3966, from which was establishedthe first of the names involved. The labels ofWright’s herbarium refer only to the countrybut do not specify the exact location in whichthey were collected (Howard 1988). However,a specimen in the Gray Herbarium, GH28253,has a note written by Wright, which says:“Balestena Feby 23.” According to Gray (citedby Underwood 1905 and Howard 1988),Balestena was a cattle farm, property of JoséBlain, located at the southern base of themountains opposite Bahía Honda, Pinar delRío. The boundaries of the farm were the SierraRangel to the north, Santa Cruz to the south,the river of the same name to the east and theTaco Taco River to the west. This locality iscurrently within the municipality of SanCristóbal, province Artemisa, and is part of thebiogeographical district Sabaloense (Borhidi1996).

The name Thrinax acuminatawas initially usedby Sauvalle (1871) for the specimen Wright3966, without description, diagnosis orreference to a previous one, which makes it anomen nudum. Subsequently the namecontinued to be used in Sauvalle (1873),Gómez de la Maza (1893) and Sargent (1899),but none of them fulfilled Article 38 of theCode. Sargent in Schumann (1901) transferredit to Coccothrinax, but this combinationcontinued to constitute a nomen nudum.Beccari (1907) first used the name Coccothrinaxacuminata (based on Coccothrinax acuminataSargent 1899) as a valid name, as it wasaccompanied by a description and a typedesignation (Wright 3966), although he didnot indicate the herbarium in which the typewas deposited. In 1972, Glassman reconsideredits taxonomic validity and indicated that, inhis opinion, the holotype was deposited in A(herbarium of the Arnold Arboretum, HarvardUniversity).

The type material used for the description thatvalidated Coccothrinax acuminata also meritscareful analysis. The specimen chosen as thelectotype is K000462859 (http://specimens.kew.org/herbarium/K000462859) deposited at Kew(Fig. 3), because Beccari used the material for


45

his description of the reproductive charactersand handwrote an annotation, “Coccothrinaxacuminata (Sargent) O. Becc III/1907.” Thissame specimen was examined previously byJ.D. Hooker, who handwrote on the sheet,“Thrinax acuminata Griseb. & Wendl. Cuba7/[18]71,” apparently referring to the nomennudum used by Sauvalle in 1871. This specimenat Kew, with Beccari’s handwritten annotation,must be the lectotype of C. acuminata. Thissupersedes all other type designations, such as“isotype of Coccothrinax acuminata” on thespecimens of Wright 3966 held at GH (GrayHerbarium of Harvard University). The claimby S.F. Glassman (1972) that the type is aspecimen deposited in A is in error; thespecimens are present only in GH (GH28253,GH28254 and GH28255) and not in otherherbaria of Harvard. Read in 1969 noted asisotypes, with corresponding annotations,duplicates of Wright 3966 in F, NY and US,while Kellogg annotated those of GH.However, none of them had been indicated as

types in any publications. In this article, a totalof 19 new isolectotypes are indicated, amongwhich, in addition to those previouslymentioned, are those existing in BRU, FI, GH,NY, P and US.

In later years, Beccari (1913), Burret (1929) andDahlgren (1936) accepted the nameCoccothrinax acuminata, but León (1939)considered it to be synonymous with C.miraguama (Kunth) Becc. var. miraguama.Many authors (León 1946, Muñiz & Borhidi1982, Henderson, Galeano & Bernal 1995,Moya & Leiva 2000, Govaerts & Dransfield,2005, Govaerts et al. 2011, Acevedo Rodríguez& Strong 2012, Greuter & Rankin, 2016)followed León (1939).

On the other hand, Beccari (1913) used thedesignation Coccothrinax miraguano (novo-geronensis), without specifying infraspecificrank, for flower and seed drawings of thespecimen Curtiss 423, although he did notclarify the herbarium he consulted. It isassumed that the specimen is present in hisherbarium at FI (Natural History Museum,Florence, Italy). He accompanied all this with


46

3. Kew specimen of Wright 3966 with Beccari’shandwritten annotation (barcode K 000462859),selected here as lectotype of Coccothrinax acuminataBecc. © copyright of the Board of Trustees of theRoyal Botanic Gardens, Kew.

4. Coccothrinax miraguama var. novo-geronensis Becc.A. Curtiss 423 with Beccari’s handwritten annotation(barcode FI 051884), selected here as lectotype. B.Original drawings for Figure 169 in Beccari (1913).© copyright of the Natural History Museum ofFlorence.

A

B

a diagnosis in which he pointed to the “almostsessile” flowers of this palm from the Isla dePinos (currently Isla de la Juventud) as thefeature differentiating it from C. miraguanofrom mainland Cuba. Beccari (1931) tran-scribed this name as a variety.

León (1939), although he mentioned Beccari(1913), was unaware of Dahlgren’s (1936)designation and indicated Curtiis 423 as oneof the paratypes of C. miraguama var. arenicolaLeón, described in his work. Glassman (1972)accepted Dahlgren (1936) and indicated theduplicate deposited in US as holotype, acriterion not shared by the authors of thisarticle, as explained later. Henderson, Galeanoand Bernal (1995) considered Coccothrinaxmiraguama var. novo-geronensis Becc. as asynonym of Coccothrinax miraguama, but thetaxon has been ignored by all the followingauthors: Burret (1929), León (1946), Muñiz &Borhidi (1982), Moya and Leiva (2000),Govaerts and Dransfield (2005) and Govaertset al. (2011), Acevedo Rodríguez and Strong(2012) and Greuter and Rankin (2016).

To designate the lectotype of Coccothrinaxmiraguano var novo-geronensis, priority wasgiven to samples deposited in FI, where Beccaricarefully studied Coccothrinax materials fromthe West Indies. The specimen FI 051884 isdesignated as a lectotype, which Beccari (1913)used for the diagnosis and drawings (Fig. 4).The statement made by Glassman (1972) isrejected, because there is no evidence that the

US duplicates were reviewed by Beccari;moreover, to be a holotype, the specimenwould have to have cited explicitly by Beccari,which it was not. None of the 17 duplicates ofCurtiss 423, deposited in CM, FI, HAC, K, LE,M, MO, NY, and US, had been previouslymentioned as part of the typological materialof Coccothrinax miraguama var. novo-geronensisBecc.

Finally, Coccothrinax miraguama var. arenicolawas described by León (1939), who quotes inthe protologue three specimens from the samelocality: León 16146, León 16147 and León16148, without designating a holotype. Leónhimself (1946) named it as a variety,nevertheless Borhidi and Muñiz (1971)changed the rank to subspecies. Subsequentauthors have followed three different points ofview: 1) Those who continued to consider itas a variety (Glassman, 1972 and AcevedoRodríguez & Strong, 2012). 2) Those whoplaced it in the subspecies rank (Muñiz &Borhidi 1982, Moya & Leiva 2000, Govaerts &Dransfield 2005, Govaerts et al. 2011, Greuter& Rankin 2016). 3) Those who did notrecognize its taxonomic validity andconsidered it as a synonym of Coccothrinaxmiraguama (Kunth) Becc. (Henderson, Galeano& Bernal 1995).

To designate the lectotype for Cocco-thrinax miraguama var. arenicola, priority wasgiven to the materials cited in the protologuebelonging to LS (currently in HAC), as it is theherbarium where Brother Leon worked duringhis entire professional stay in Cuba. Itdesignated as lectotype León 16146[HACLS4387], on whose label León wrote“type.” The numbers: León 16147 [HAC-LS4386], used to describe flowers and León16148 [HAC-LS4385], remain only asparatypes. None of the seven duplicates of León16146, in HAC-UO and US, had beenpreviously been mentioned as part of the typematerial of Coccothrinax miraguama subsp.arenicola (León) Borhidi & O.Muñiz.

According to the above, the proposednomenclature for the taxa analyzed is asfollows:

Coccothrinax acuminata Becc., Webbia. 2:313 (1907). Thrinax acuminata Griseb. & H.Wendl., in Sauvalle, Anales Acad. Ci. Med.Habana, 8: 563 (1871), nom. nud. Type.CUBA. [Provincia Artemisa, municipio SanCristóbal], Balestena. ft., 23. Feb. [1862 or1864]. Wright 3966 (lectotype, heredesignated, K 000462859 [photo!];


47

5. Vegetation at Los Indios, Isla de la Juventud, withCoccothrinax acuminata. Photo by Michael Calonje.

isolectotypes, here designated: BRU 55644[photo!], BRU 55645 [photo!], V 92098F1[photo!], V 92098F2 [photo!], V 92098F3[photo!], FI 51879 ex K [photo!], GH 28253[photo!], GH 28254 [photo!], GH 28255[photo!], K 462858 [photo!], NY 73060[photo!], NY 73076 [photo!], NY 73077[photo!], NY 73078 [photo!], NY 73079[photo!], P 725688 [photo!], P 725689[photo!], US 87368 [photo!], US 87369[photo!]).

Coccothrinax miraguama var. novo-geronensisBecc., Ann. Roy. Bot. Gard. Calcutta 13: 336(1931). Coccothrinax miraguama [withoutrank] (novo-geronensis) Becc., Pomona Coll. J.Econ. Bot. 3: 409 (1913), synon. nov. Type.CUBA. [Municipio Isla de la Juventud], dryground of poor quality near Nueva Gerona,Isla de Pinos, W.I., April [ft.]-1 May [fl.] 1904,Curtiss 423 (lectotype, here designated, FI051884 [photo!]; isolectotypes, heredesignated: CM 422028 [photo!], CM422029 [photo!], G 305367 [n.v.], K 632580[photo!], K 632581 [photo!], LE 793 [photo!],HAC!, M 208181 [photo!], MO 559592 [n.v.],MO 559593 [n.v.], NY 1661902 [photo!], NY1662094 [photo!], NY 1662095 [photo!], NY1662105 [photo!], US 14965 [photo!],VT117062 [photo!], MO 559592 [n.v.], MO559593 [n.v.]).

Coccothrinax miraguama subsp. arenicola(León) Borhidi & O.Muñiz, Bot. Közlem. 58:175 (1971). Coccothrinax miraguama var.arenicola León, Mem. Soc. Cub. Hist. Nat.“Felipe Poey’” 13: 114 (1939), synom. nov.Type: CUBA. [Provincia Pinar del Río,municipio Guane], sabana arenosa, haciendaSabanalamar, El Sábalo (Pinar del Río), ft., 20.Aug. 1934, León 16146 (lectotype, heredesignated, HAC-LS4387!; isolectotypes, heredesignated: HAC-UO1!, HAC-UO2! HAC-UO3!, US14992 [photo!], US14993 [photo!],US14994 [photo!], US14995 [photo!]).

Specimens examined: CUBA. Other specimensexamined to which the identification isupdated as Coccothrinax acuminata. Thenumber of duplicates of each specimen in thesame herbarium, is indicated in brackets.

Palmer 877 US [photo!]; Shafer 299 HAC!, CM[photo! 3×], NY [photo! 2×]; Baker 4808 HAC!,FI [photo!]; Hermann 587NY [photo!]; Hermann714HAC!, FI [photo!]; Hermann 839HAC! [2×],FI [photo!]; Jennings 156 NY [photo!], CM[photo! 2×]; Jennings 623 CM photo!]; Britton6652 NY [photo! 2×], US [photo!]; Britton 9748NY [photo! 2×], US [photo!]; Britton 10089 NY

[photo! 2×]; Britton 14227 CM [photo!], NY[photo!], US [photo! 2×]; Shafer 10561 NY[photo! 2×], US [photo!]; Shafer 10921 NY[photo! 2×], US [photo! 2×]; Hermann 7909,HAC! [5×]; Hermann s.n.HAC! [2×]; León 16147HAC!; León 16148 HAC!; León 16150 HAC!;León 17034 HAC! [3×], US [photo! 2×]; León17035 HAC!, US [photo! 2×]; León 17466HAC!;León 17467 HAC!; León 18596 HAC!); León18734HAC!; León 18851HAC!; Killip 32239US[photo! 2×]; Acuña 19849 HAC! [2×]; Alain6913 HAC!; Verdecia RV12/23 HMC [photo!2×]; Verdecia RV12/24 HMC [photo! 3×];Verdecia RV12/28HMC [photo! 3×], NY [photo!4×]; Verdecia RV12/35 HMC [photo! 2×], NY[photo! 4×].

Notes: The species Coccothrinax acuminatabelongs to subsection Coccothrinax of sectionCoccothrinax, according to the classification ofMuñiz and Borhidi (1982), and Miraguamacomplex of the Pauciramosa Group, accordingto the informal classification of Nauman andSanders (1991).

Distribution: Provinces Artemisa, municipalitySan Cristóbal; and Pinar del Río, municipalitiesConsolación del Sur, Guane, Mantua, Pinar delRío, Sandino, San Juan y Martínez y San Luis,(León 1939, Urquiola et al. 2001), andmunicipality Isla de la Juventud (León 1939).

Biogeography: Western Cuba subprovince,district Guanahacabibense (sector Peninsu-laricum), districts Geronense, Indionense, Pinar-ense and Sabaloense (sector Pinaricum) anddistrict Viñalense (sector Rosaricum) (Borhidi1996).

Habitat: The species grows in secondarysavannas, seminatural savannas, coastal andsubcoastal thorny shrublands and pine forest,on sandy-quartzite substrate, mainly on whitesandy soils (Fig. 5 & Front Cover), alluvial soilsand rarely on limestone.

Vernacular names: Guanito and miraguano (León1939), yuraguana (González-Oliva et al. 2015).

Conservation status: Least Concern (LC) sensuGonzález-Torres et al. (2016) for thecategorization of Coccothrinax miraguamasubsp. arenicola. According to González-Olivaet al. (2015), present in GuanahacabibesNational Park, in the Los Indios and LosPretiles Ecological Reserves, in the Sierra deContadores-Cayo Ratones and San Ubaldo-Sabanalamar Managed Floristic Reserves, inthe La Cañada Managed Resource ProtectedAreas and in the Península de GunahacabibesBiosphere Reserve. León (1939) reported


48

damage to populations by periodic fire andcutting by the campesinos, who use the leaves.González-Oliva et al. (2015) reported that it isused for rustic constructions.

Acknowledgments

We are grateful to Drs. Scott Zona and JohnDransfield, who critically reviewed themanuscript. We acknowledge Paul Craft forhis help and logistic contribution to the mainauthor during fieldwork in the Isla de laJuventud and Pinar del Río, also for thetranslation into English. We thank Chiara Nepiof FI for sending us “The Palms of OdoardoBeccari.” We thank Meghann S. Tonner andIngrid Lin of US, Bonnie L. Isaac of CM, WalterKittredge of A, GH, Chiara Nepi and EgildoLuccioli of FI, Tim Whifeld of BRU, andBanessa Falcón (LE and M) for makingspecimens available for study, to Dr. RosaRankin, from the National Botanic Garden ofLa Habana, for their advice regarding theapplication of certain articles of the ICN, andto Rafael Govaerts for his critical expertopinions on nomenclature.

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